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Pollinating Fig Wasp Ceratosolen Solmsi Adjusts the Offspring Sex

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Pollinating Fig Wasp Ceratosolen Solmsi Adjusts the Offspring Sex Insect Science (2013) 20, 228–234, DOI 10.1111/j.1744-7917.2011.01495.x ORIGINAL ARTICLE Pollinating fig wasp Ceratosolen solmsi adjusts the offspring sex ratio to other foundresses Hao-Yuan Hu1, Zhong-Zheng Chen1, Zi-Feng Jiang2, Da-Wei Huang3,4, Li-Ming Niu5 and Yue-Guan Fu5 1Key Laboratory of Biotic Environment and Ecological Safety in Anhui Province, College of Life Sciences, Anhui Normal University, Wuhu, Anhui Province, China, 2Department of Biology, University of Maryland, College Park, Maryland, USA, 3Key Laboratory of Zoological Systematics and Evolution, Institute of Zoology, Chinese Academy of Sciences, Beijing, China, 4Plant Protection College, Shandong Agricultural University, Tai’an, Shandong Province, China, 5Environment and Plant Protection Institute, Chinese Academy of Tropical Agricultural Sciences, Danzhou, Hainan Province, China Abstract Local mate competition theory predicts that offspring sex ratio in pollinating fig wasps is female-biased when there is only one foundress, and increased foundress density results in increased offspring sex ratio. Information of other foundresses and clutch size have been suggested to be the main proximate explanations for sex ratio adjustment under local mate competition. Our focus was to show the mechanism of sex ratio adjustment in a pollinating fig wasp, Ceratosolen solmsi Mayr, an obligate pollinator of the functionally dioecious fig, Ficus hispida Linn., with controlled experiments in the field. First, we obtained offspring from one pollinator and offspring at different oviposition sequences, and found that offspring sex ratio decreased with clutch size, and pollinators produced most of their male offspring at the start of bouts, followed by mostly females. Second, we found that offspring sex ratio increased with foundress density, and pollinators did adjust their offspring sex ratio to other females in the oviposition patches. We suggest that when oviposition sites are not limited, pollinators will mainly adjust their offspring sex ratio to other foundresses independent of clutch size changes, whereas adjusting clutch size may be used to adjust sex ratio when oviposition sites are limited. Key words behavior, co-evolution, local mate competition, mutualism, pollinating fig wasp, sex ratio adjustment Introduction highly structured population, such as mating only between full siblings, evolutionary stable strategy models predict a Sex ratio in a local environment has been taken as a model female-biased sex ratio (Hamilton, 1967; Maynard Smith, of sex ratio theory (Hamilton, 1967; Frank, 1985; Herre, 1976). In haplodiploid hymenoptera, with haploid males 1985, 1987; West et al., 2000; Steiner & Ruther, 2009; (developed from unfertilized eggs) and diploid females West, 2009) and a case study of adaptation in evolution- (developed from fertilized eggs), the primary sex ratio is ary biology (Fellowes et al., 1999). In a random-mating determined by the mothers’ control of fertilization at the populations of infinite size, natural selection favors equal time of oviposition (Charnov, 1982). investment in both sexes, and an equal sex ratio is an There may be natural selection on foundresses to adjust evolutionary stable result (Fisher, 1930). However, in a their offspring sex ratio because of local mate competition (LMC) (Charnov, 1982; West et al., 2005). When patches are founded by one female and all mating occurs between Correspondence: Hao-Yuan Hu, College of Life Sciences, siblings, brothers may compete locally with each other Anhui Normal University, Wuhu, Anhui Province 241000, for mating with their sisters. Such LMC selects for fewer China. email: [email protected] male offspring (Sheldon & West, 2002). If offspring in a C 2012 The Authors 228 Insect Science C 2012 Institute of Zoology, Chinese Academy of Sciences Sex ratio adjustment of a fig wasp 229 patch originate from two foundresses, the value of sons Academy of Tropical Agricultural Sciences (CATAS), increases because sons have the potential to mate with the Danzhou, Hainan province, China (19◦30.410S, daughters of other foundresses, and a less female-biased 109◦29.340E). Hainan Island is located south of the offspring sex ratio is preferred (Hamilton, 1967, 1979; China mainland separated by the Qiongzhou Strait, ap- Herre, 1985, 1987; Werren, 1987). Sex ratios increase proximately 40 km wide. The climate includes well- with foundress number, as LMC models predict, in many defined dry (November–April) and rainy (May–October) parasitoid wasps (reviewed in King, 1993) and some fig seasons, and the annual mean temperature is 24.3◦C, with wasps (e.g., Raja et al., 2008) the lowest mean temperature in February (18.2◦C) and the The proximate explanations for such sex ratio ad- highest in July (29.6◦C). justment under LMC may differ among species (Moore Ficus hispida is a functionally dioecious and free- et al., 2005). Some reports have argued that clutch size standing tree, with inter-sexually synchronously flowering might be one of the main explanations when more par- phenologies. Most fruits grow on fruit branches, and can asitoid wasps or fig wasps oviposit in a limited patch be kept away from pollinators using specially made se- (Kjellberg et al., 2005; Moore et al., 2005; Raja et al., curing nylon bags during fig receptivity. Pollinators can 2008). When multiple foundresses oviposit in figs, ovipo- be collected from mature male fruits in mesh-lidded pots sition site limitation would lead to reduction of offspring just before wasp emergence (Hu et al., 2009, 2010). reproduction (Moore & Greeff, 2003). Limited clutch size and the oviposition order of fertilized and unfertilized eggs may lead to a less female-biased sex ratio (Kjellberg Offspring sex ratio and clutch size et al., 2005; Moore et al., 2005; Raja et al., 2008; Sun et al., 2009). Another explanation of sex ratio adjustment To investigate the effect of clutch size on offspring sex under LMC is foundress density affecting probability of ratio, we obtained offspring from one-foundress figs. Fe- ovipositing a son and that probability being independent male pollinators were collected as above and immedi- of order. Foundresses may adjust their offspring sex ratios ately introduced into the protected receptive figs with a by the information of other foundresses or of oviposi- soft brush. One pollinator was allowed to enter each fig, tion in a patch left by other foundresses (Hamilton, 1967, and then the figs were re-bagged individually; 241 one- 1979; Herre, 1985, 1987; Kinoshita et al., 2002; Moore foundress figs were obtained from seven male F. hispida et al., 2002; Pereira & Prado, 2006; Herre et al., 2008; tress from January to December in 2007. Abe et al., 2009; Somjee et al., 2011). Small clutch size can be obtained by ceasing oviposi- In some pollinating fig wasps, clutch size would be tion after the pollinator’s entry. Pollinators oviposit inside the major explanation to sex ratio adjustment under LMC figs, and it is hard to cease the oviposition and keep devel- (Kjellberg et al., 2005; Moore et al., 2005; Raja et al., opment of the figs at the same time. Recent research has 2008; Sun et al., 2009). The effect of sex adjustment to used poisons such as insecticide pyrethrum (Raja et al., other foundresses has been called in to question (Kjellberg 2008) and ether (Sun et al., 2009) to cease the oviposition. et al., 2005). But oviposition sites would not be always Patino et al. (1994) showed that when figs reach temper- limited when only several foundresses lay eggs in a patch. atures higher than 8◦C above the ambient temperature When oviposition sites in a patch are sufficient, clutch size in full sunlight, pollinators inside figs would be killed. should be unaffected by foundress density. Given this, we Warm water has been used to kill pollinators inside figs predict that the effect of clutch size on sex ratio adjustment (Hu et al., 2009). In this article, warm water (36–40◦C), should be weak when oviposition sites are rarely limited, 8◦C above ambient temperature, was injected into a plas- as in some figs with plenty of florets, where oviposition tic bag (≈ 2 L), and the figs were put into the bag to sites are always enough. We test this with Ceratosolen kill pollinators in situ. One-foundress figs were opened at solmsi Mayr, an obligate pollinator of the functionally 2-min intervals after the figs were put into warm water to dioecious fig, Ficus hispida Linn. examine whether the foundress was dead or not. Ten min- utes were selected because foundresses inside large and small figs were all dead then. Twenty one-foundress figs Materials and methods warmed at 4 h after pollinator entry were protected from entry by other foundresses with nylon bags, and 13 treated Study species and site figs matured normally. Previous research has shown that C. solmsi could only live in figs for less than 24 h, and Experiments on Ficus hispida Linn. – Ceratosolen the duration of oviposition lasts less than 9 h (Yang et al., solmsi Mayr mutualism were conducted at the Chinese 2002). Thus, three treatments were designed: one without C 2012 The Authors Insect Science C 2012 Institute of Zoology, Chinese Academy of Sciences, 20, 228–234 230 H. Y. Hu et al. warming, the other two with 2- and 4-h intervals (from pollinator entering to warming) respectively. More than 50 one-foundress figs were used in each treatment on two male F. hispida trees in September 2007 and August 2008, respectively. Offspring sex ratio and foundress density To investigate the effect of foundress density on the offspring sex ratio, the following experiments were conducted. Five treatments were designed with 1–5 foundresses respectively, and more than 30 figs were used in each treatment. Female pollinators were collected as above and immediately introduced into the protected re- ceptive figs. Wasps introduced to multiple-foundress figs Fig. 1 Correlation between clutch size and sex ratio of Cer- were from mature figs on different fig trees, which en- atosolen solmsi in one-foundress figs.
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